Scot E. Hagerthey

MULTIPLE REGIME SHIFTS IN A SUBTROPICAL PEATLAND: COMMUNITY-SPECIFIC THRESHOLDS TO EUTROPHICATION

Ecosystems have a natural resilience to perturbations, where resilience is the magnitude of a disturbance that an ecosystem can resist before changes in structure, function, and services result in a regime shift. The Everglades region of Florida, USA, has been detrimentally impacted by phosphorus (P) enrichment and a regime shift from Cladium (sawgrass) to Typha (cattail) marsh has been described. We examine another facet of the low nutrient Everglades stability regime, open-water sloughs, to determine if eutrophication leads to similar regime shifts. We analyzed surface water P and soil P as controlling variables that, once a critical threshold is surpassed, alter ecosystem state variables. Nonlinear relationships between P and vegetation were observed along a northern Everglades eutrophication gradient. In addition to the Cladium-Typha regime shift, a second independent regime shift, slough-Typha, was identified. Synoptic surveys of 49 sloughs within the boundary between the slough and Typha regime revealed that surface water total phosphorus (TP) and the benthic algal floe layer (BAFL) were the controlling variables, with critical thresholds of 11 ug/L and 412 mg/kg, respectively. The slough regime below these thresholds was characterized by calcareous periphyton (BAFL TP = 298 mg P/kg; BAFL calcium = 149 g Ca/kg). Above the TP thresholds, vegetation composition shifted to open-marsh species with significantly higher BAFL TP (700 mg P/kg) and total organic carbon (TOC) (350 g C/kg). A second BAFL TP threshold occurred at 712 mg P/kg, above which Nymphaea dominated and BAFL TP (1034 mg P/kg) and TOC (417 g C/kg) significantly increased. Nymphaea sloughs transitioned to the Typha regime. The boundary reflects the loss of ecosystem resilience due to eutrophication. Both low-nutrient stability regimes (slough and Cladium) lie precariously close to the P critical threshold but differ in how eutrophication is absorbed and resisted. The slough regime transitions rapidly through a series of ecosystem state changes linked to positive feedback loops that affect P dynamics, whereas the Cladium regime does not. An adaptive management strategy has been implemented to address the surface water TP threshold; however, to ensure successful restoration of the Everglades, the BAFL and soil TP thresholds also need to be considered.

Landscape Patterns of Periphyton in the Florida Everglades

Periphyton is an abundant and ubiquitous feature of the Florida Everglades, often forming thick mats that blanket shallow sediments and submersed plants. They are considered to be primary ecosystem engineers in the Everglades by forming and stabilizing soils, controlling concentrations of nutrients and gases, and supplying food and structure for other organisms. Distribution patterns are related to underlying physicochemical gradients as well as those hydrologic changes imposed by water management. Because communities respond rapidly to environmental change, their use has been advocated to provide indication of system degradation or restoration. The authors review studies on the distribution of periphyton in the Everglades, highlighting major findings relevant to water management, and also areas where additional exploration is necessary.

Recent and historic drivers of landscape change in the everglades ridge, Slough, and Tree Island Mosaic

More than half of the original Everglades extent formed a patterned peat mosaic of elevated ridges, lower and more open sloughs, and tree islands aligned parallel to the dominant flow direction. This ecologically important landscape structure remained in a dynamic equilibrium for millennia prior to rapid degradation over the past century in response to human manipulation of the hydrologic system. Restoration of the patterned landscape structure is one of the primary objectives of the Everglades restoration effort. Recent research has revealed that three main drivers regulated feedbacks that initiated and maintained landscape structure: the spatial and temporal distribution of surface water depths, surface and subsurface flow, and phosphorus supply. Causes of recent degradation include but are not limited to perturbations to these historically important controls; shifts in mineral and sulfate supply may have also contributed to degradation. Restoring predrainage hydrologic conditions will likely preserve remaining landscape pattern structure, provided a sufficient supply of surface water with low nutrient and low total dissolved solids content exists to maintain a rainfall-driven water chemistry. However, because of hysteresis in landscape evolution trajectories, restoration of areas with a fully degraded landscape could require additional human intervention.

Evaluation of pigment extraction methods and a recommended protocol for periphyton chlorophyll a determination and chemotaxonomic assessment

Many methods have been proposed to extract and quantify algal pigments. Comparative studies have found that pigment extraction efficiency varies among solvent and mechanical disruption protocols due to differential cellular resistance, thereby, leading to potential misinterpretation of pigment data. When the type or resistance of algae are unknown, a method is required that efficiently extract pigments from all taxonomic groups. The objective of this study was to develop a simple and efficient one stage periphyton pigment extraction protocol by comparing the extractability of four solvents (acetone, methanol, methanol/acetone, and methanol/acetone/N,N‐dimethylformamide), the effects of grinding, and the effects of freeze‐drying. The best overall extraction was obtained using freeze‐dried samples extracted with methanol/acetone/DMF/water (MAD). Eighty‐six percent more chlorophyll was extracted when the sample was freeze‐dried relative to fresh/frozen samples extracted with 90% acetone. Freeze‐drying greatly improved the extraction of both polar and non‐polar (lipophilic/hydrophobic) pigments while MAD increased the extractability of polar pigments and improved peak resolution of all pigments. Chemotaxonomic assessment differed between samples that were fresh/frozen or freeze‐dried before extraction. The relative abundance of cyanobacteria was greater for freeze‐dried material compared with fresh/frozen due to the improved extractability of cyanobacterial pigments. Based on the results of this study, the traditional approach of 90% acetone as a solvent is not recommended for periphyton samples containing cyanobacteria or when the composition of the mat is unknown. The combination of freeze‐drying and MAD was sufficient for the extraction of pigments from a periphyton mat containing filamentous cyanobacteria, green algae, and diatoms.

Everglades Periphyton: A Biogeochemical Perspective

Periphyton is an important component of the Everglades biogeochemical cycle but remains poorly understood. From a biogeochemical perspective, periphyton is a dense aggregation of diverse microorganisms (autotrophic and heterotrophic) and particles (mineral and detrital) imbedded within an extracellular matrix. The authors synthesize Everglades periphyton biogeochemistry and diversity at the ecosystem and community scales. The primary regulator of biogeochemical processes (material flux, transformation, and storage) is photosynthesis, which controls oxidation-reduction potentials and heterotrophic metabolism. Eutrophication and hydrologic alterations have resulted in fundamental periphyton biogeochemical differences. Elucidation of these processes is required to predict and interpret responses to ecosystem restoration.

Composition of extracellular polymeric substances from periphyton assemblages in the Florida Everglades.

In wetland habitats, periphyton is a common component of open‐water areas with species assemblage determined by local water quality. Extracellular polymeric substances (EPS) secreted by algae and bacteria give structure to periphyton, and differences in EPS chemistry affect the functional roles of these polymers. The Florida Everglades provide a unique opportunity to study compositional differences of EPS from distinctive algal assemblages that characterize areas of differing water chemistry. Water conservation area (WCA)‐1 is a soft‐water impoundment; periphyton was loosely associated with Utricularia stems and amorphous in structure, with a diverse desmid and diatom assemblage, and varying cyanobacterial abundance. Extracellular polymers were abundant and were loosely cell‐associated sheaths and slime layers in addition to tightly cell‐associated capsules. The EPS were complex heteropolysaccharides with significant saccharide residues of glucose, xylose, arabinose, and fucose. Carboxylic acids were also prominent, while ester sulfates and proteins were small components. Structured, cohesive cyanobacteria‐dominated periphyton was observed in WCA‐2A, a minerotrophic impoundment, and filaments were heavily encrusted with calcium carbonate and detrital matter. EPS were primarily cell‐associated sheaths, and polymer residues were dominated by glucose, xylose, fucose, and galactose, with uronic acids also a significant component of the polymers. Principal components analysis revealed that periphyton community assemblage determined the monosaccharide composition of EPS, which ultimately determines a range of biogeochemical processes within the periphyton.

Water Conservation Area 1: A Case Study of Hydrology, Nutrient, and Mineral Influences on Biogeochemical Processes

At the northern tip of the remnant Everglades, Water Conservation Area 1 is the only remaining softwater peatland in the ecosystem. The spatial pattern of altered hydrology, anthropogenic nutrient, and mineral enrichment is distinct, with biogeochemical processes driven by a north-south hydrologic gradient combined with west-east nutrient and mineral gradients. Hydrology effects on carbon cycling are evident by the 10–20% lower average soil carbon concentrations in the drier oxidizing regions of the north, compared with the ponded environment in the south. Elevated nutrient and mineral inputs also increase carbon loss by causing changes in species composition, substrate quality, and microbial activity. Water management may be optimized to limit mineral intrusion and peat oxidation, while also meeting water depth requirements for habitat and wildlife, such that ecological tradeoffs are minimized.

PRESENCE AND DIVERSITY OF ALGAL TOXINS IN SUBTROPICAL PEATLAND PERIPHYTON: THE FLORIDA EVERGLADES, USA1

Production of toxic secondary metabolites by cyanobacteria, collectively referred to as cyanotoxins, has been well described for eutrophied water bodies around the world. However, cohesive cyanobacterial mats also comprise a significant amount of biomass in subtropical oligotrophic wetlands. As these habitats generally do not support much secondary production, cyanotoxins, coupled with other physiological attributes of cyanobacteria, may be contributing to the minimized consumer biomass. Periphyton from the Florida Everglades has a diverse and abundant cyanobacterial assemblage whose species produce toxic metabolites; therefore, by screening periphyton representative of the greater Everglades ecosystem, six different cyanotoxins and one toxin (domoic acid) produced by diatoms were identified, ranging in content from 3 × 10−9 to 1.3 × 10−6 (g · g−1), with saxitoxin, microcystin, and anatoxin‐a being the most common. While content of toxins were generally low, when coupled with the tremendous periphyton biomass (3–3,000 g · m−2), a significant amount of cyanotoxins may be present. While the direct effects of the toxins identified here on the local grazing community need to be determined, the screening process utilized proved effective in showing the broad potential of periphyton to produce a variety of toxins.